Interactions of nutrient and water availability control growth and diversity effects in a Salix two-species mixture

Plant growth is constrained by resource availability and interactions among limiting resources—abundance in one resource (e.g., nutrients) might promote growth, thereby causing the depletion of other resources (e.g., water), potentially inducing stress or mortality. In a diverse plant community, complementary resource use has been hypothesized to increase the overall productivity, but how diversity effects vary with interacting water and nutrient limitation and through time is not known. Here, we address this knowledge gap in a controlled pot experiment where species composition (two Salix species in monoculture or mixture), nutrient addition, and watering frequency (for fixed total water inputs) were varied during two growing seasons. High nutrient availability promoted plant growth and nitrogen accumulation at the pot scale, as well as increased allocation aboveground, but also triggered more intense water stress and mortality, as larger plants depleted soil water during warm periods. Supplying water more frequently slightly alleviated water stress under high nutrient availability, thus promoting growth and nitrogen accumulation. The species mixtures performed better than the average of the mixture constituents (positive net diversity effects) and increasingly so through time. The complementarity and selection effects, respectively, increased and decreased under both high nutrient availability and high watering frequency. Overall, these results suggest that as plants grow larger, plant interactions and resource partitioning intensify, causing the positive diversity effects, but also that drought consequences might be exacerbated in plant communities rapidly growing thanks to high nutrient supply

Functional traits of individual varieties as determinants of growth and nitrogen use patterns in mixed stands of willow (Salix spp.)

Short rotation plantations of willows (Salix spp.) have high biomass production potential in many parts of the world, and may frequently support ecosystem services related to nutrient cycling. A plantation management enhancing favorable environmental impacts that are conducive to maintaining ecosystem services is a main challenge in establishing sustainable biomass production systems. There is evidence supporting the hypothesis that biomass production and nutrient cycling can be increased by supporting ecosystem niche differentiation (complementarity) through enhancing the number of plant species or varieties grown in the stand. However, the specific trait values of the individual components (e.g., varieties) in a mixed community could also be more important than the community diversity per se. We assessed, at community level, the plant trait profiles related to growth and nitrogen (N) use for four different Salix varieties that were taxonomically distinct at species or genotype level ('Bjorn', 'Jorr', 'Loden', 'Tora') and field-grown in unfertilized plots of pure and mixed commu-nities during one cutting cycle in Central Sweden. The aims were to use elements of functional growth analysis for exploring the mechanistic relationships between various traits related to growth and N use at stand level in our pure and mixed willow communities; and to address two hypotheses related to (i) the effect of diversity level on above-ground traits linked to growth, N uptake efficiency, N productivity and N conservation; and (ii) the influence of individual variety identities on the growth and N use traits observed in a mixture. Diversity level had no significant effect on the traits assessed here, and we thus found no evidence in support of our hypothesis that traits linked to growth, N uptake and use are significantly affected by the diversity level per se. In most but not all cases, the admixing effects on trait values were explained by the effects of the individual variety characteristics assessed in monocultures in combination with their relative share in the respective mixtures. The absence or presence of individual varieties strongly affected community-averaged (stand level) trait values. Therefore, the design of desirable variety mixtures is suggested that combine, for example, the high nutrient conversion efficiency that certain varieties achieve in mixed stands with the specific nutrient acquisition characteristics of other varieties

Site-dependent relationships between fungal community composition, plant genotypic diversity and environmental drivers in a Salix biomass system

Soil fungi are strongly affected by plant species or genotypes since plants modify their surrounding environment, but the effects of plant genotype diversity on fungal diversity and function have not been extensively studied. The interactive responses of fungal community composition to plant genotypic diversity and environmental drivers were investigated in Salix biomass systems, posing questions about: (1) How fungal diversity varies as a function of plant genotype diversity; (2) If plant genotype identity is a strong driver of fungal community composition also in plant mixtures; (3) How the fungal communities change through time (seasonally and interannually)?; and (4) Will the proportion of ECM fungi increase over the rotation? Soil samples were collected over 4 years, starting preplanting from two Salix field trials, including four genotypes with contrasting phenology and functional traits, and genotypes were grown in all possible combinations (four genotypes in Uppsala, Sweden, two in Rostock, Germany). Fungal communities were identified, using Pacific Biosciences sequencing of fungal ITS2 amplicons. We found some site-dependent relationships between fungal community composition and genotype or diversity level, and site accounted for the largest part of the variation in fungal community composition. Rostock had a more homogenous community structure, with significant effects of genotype, diversity level, and the presence of one genotype (“Loden”) on fungal community composition. Soil properties and plant and litter traits contributed to explaining the variation in fungal species composition. The within-season variation in composition was of a similar magnitude to the year-to-year variation. The proportion of ECM fungi increased over time irrespective of plant genotype diversity, and, in Uppsala, the 4-mixture showed a weaker response than other combinations. Species richness was generally higher in Uppsala compared with that in Rostock and increased over time, but did not increase with plant genotype diversity. This significant site-specificity underlines the need for consideration of diverse sites to draw general conclusions of temporal variations and functioning of fungal communities. A significant increase in ECM colonization of soil under the pioneer tree Salix on agricultural soils was evident and points to changed litter decomposition and soil carbon dynamics during Salix growth

Biodiversity-ecosystem function in a willow biomass production system

The relationship between biodiversity and ecosystem function is an important issue in ecology. Stands of Salix (willow) are suitable model systems to study this relationship. Salix and other crops are mainly grown in monoculture in agricultural systems. Species or genotypes grown in monoculture share the same functional traits and can therefore be expected to compete strongly for resources. In contrast, different species or genotypes grown in mixed culture vary in their functional traits and may use different niches, leading to reduced competition. Thus, higher diversity in functional traits can increase ecosystem functions such as productivity and litter decomposition. This thesis examined how individual Salix genotypes affect community shoot biomass, litter decomposition and fungal diversity when grown in pure and mixed cultures of different genotypes. Three field sites were established in Central and Northern Europe (Freiburg and Rostock in Germany, Uppsala in Sweden). Within each location, plots were planted with pure and mixed communities of four Salix genotypes (‘Björn’, ‘Jorr’, ‘Loden’ and ‘Tora’) that differ in their morphological and functional traits. In addition to the field study, the two taxonomically and physiologically most distinct genotypes (‘Loden’ and ‘Tora’) were grown under two different nutrient treatments in pure and mixed communities in a pot study. Genotypes exerted different influences on the Salix community in which they were grown, although there was no general increase in productivity, decomposition rate or fungal diversity with increasing genetic richness. Among other findings, one genotype (‘Jorr’) enhanced productivity and litter decomposition when grown in mixed communities. Another genotype (‘Tora’) had a negative effect on productivity, litter decomposition and fungal diversity when added to a community. ‘Tora’ benefited from the presence of other genotypes, but under nutrient poor conditions it performed worse in the presence of a competitor. Litter chemistry differed between genotypes, and decomposition showed a distinct pattern between sites. Fungal communities were affected by different drivers including leaf chemistry, soil properties and genotype identity, but not by genetic diversity. The results suggest that the specific functional trait combinations of individual genotypes affect their response to mixture as compared to monoculture and that the attributes of individual genotypes are more important than genetic richness for the ecosystem functions studied here

Traits to Ecosystems: The Ecological Sustainability Challenge When Developing Future Energy Crops

Today we are undertaking great efforts to improve biomass production and quality traits of energy crops. Major motivation for developing those crops is based on environmental and ecological sustainability considerations, which however often are de-coupled from the trait-based crop improvement programs. It is now time to develop appropriate methods to link crop traits to production system characteristics set by the plant and the biotic communities influencing it; and to the ecosystem processes affecting ecological sustainability. The relevant ecosystem processes involve the net productivity in terms of biomass and energy yields, the depletion of energy-demanding resources (e.g. nitrogen, N), the carbon dynamics in soil and atmosphere, and the resilience and temporal stability of the production system. In a case study, we compared aspects of N use efficiency in various varieties of an annual (spring wheat) and perennial (Salix) energy crop grown under two nutrient regimes in Sweden. For example, we found considerable variation among crops, varieties and nutrient regimes in the energy yield per plant-internal N (MJ g-1 yr-1), which would result in different N resource depletion per unit energy produced

Leaf litter quality coupled to Salix variety drives litter decomposition more than stand diversity or climate

Aims Decomposition of leaf litter is influenced by litter quality as determined by plant genotype and environment, as well as climate and soil properties. We studied these drivers of decomposition in communities ofSalixvarieties, hypothesizing that decomposition rates would increase under warmer climate, in more diverse communities, and with increasing litter quality of the individual varieties. Methods Litter from fourSalixvarieties was incubated in three field trials across a latitudinal gradient from Central to Northern Europe. Litter and stand properties were measured and used as predictors of decomposition. Results No significant site differences in remaining mass or nitrogen were found. Instead, effects of initial leaf litter quality on decomposition were stronger than climatic effects. Litter quality of individual varieties strongly affected decomposition, while increasing litter diversity did not. Conclusions Decomposition was controlled by variety identity depending on site, indicating that local soil conditions affect litter quality (and thus decomposition) more than macroclimate. In mixed communities, varieties producing fast-decomposing litter enhanced the litter decomposition of other components producing slow-decomposing litter, and vice versa. This implies that site conditions partly determine which varieties affect community-level decomposition and nutrient release

Two Salix Genotypes Differ in Productivity and Nitrogen Economy When Grown in Monoculture and Mixture

Individual plant species or genotypes often differ in their demand for nutrients; to compete in a community they must be able to acquire more nutrients (i.e., uptake efficiency) and/or use them more efficiently for biomass production than their competitors. These two mechanisms are often complementary, as there are inherent trade-offs between them. In a mixed-stand, species with contrasting nutrient use patterns interact and may use their resources to increase productivity in different ways. Under contrasting nutrient availabilities, the competitive advantages conferred by either strategy may also shift, so that the interaction between resource use strategy and resource availability ultimately determines the performance of individual genotypes in mixtures. The aim was to investigate growth and nitrogen (N) use efficiency of two willow (Salix) genotypes grown in monoculture and mixture in a fertilizer contrast. We explored the hypotheses that (1) the biomass production of at least one of the involved genotypes should be greater when grown in mixture as compared to the corresponding monoculture when nutrients are the most growth-limiting factor; and (2) the N economy of individual genotypes differs when grown in mixture compared to the corresponding monoculture. The genotypes ‘Tora’ (Salix schwerinii ×S. viminalis) and ‘Loden’ (S. dasyclados), with contrasting phenology and functional traits, were grown from cuttings in a growth container experiment under two nutrient fertilization treatments (high and low) in mono- and mixed-culture for 17 weeks. Under low nutrient level, ‘Tora’ showed a higher biomass production (aboveground biomass, leaf area productivity) and N uptake efficiency in mixture than in monoculture, whereas ‘Loden’ showed the opposite pattern. In addition, ‘Loden’ showed higher leaf N productivity but lower N uptake efficiency than ‘Tora.’ The results demonstrated that the specific functional trait combinations of individual genotypes affect their response to mixture as compared to monoculture. Plants grown in mixture as opposed to monoculture may thus increase biomass and vary in their response of N use efficiency traits. However, young plants were investigated here, and as we cannot predict mixture response in mature stands, our results need to be validated at field scale.This project was partly funded by the Swedish Energy Agency (project no. 36654-1). Partial support to SM was provided by Formas (project no. 2016-00998).Peer reviewedPeer Reviewe

Mixture of Salix Genotypes Promotes Root Colonization With Dark Septate Endophytes and Changes P Cycling in the Mycorrhizosphere

The roots of Salix spp. can be colonized by two types of mycorrhizal fungi (ectomycorrhizal and arbuscular) and furthermore by dark-septate endophytes. The fungal root colonization is affected by the plant genotype, soil properties and their interactions. However, the impact of host diversity accomplished by mixing different Salix genotypes within the site on root-associated fungi and P-mobilization in the field is not known. It can be hypothesized that mixing of genotypes with strong eco-physiological differences changes the diversity and abundance of root-associated fungi and P-mobilization in the mycorrhizosphere based on different root characteristics. To test this hypothesis, we have studied the mixture of two fundamentally eco-physiologically different Salix genotypes (S. dasyclados cv. ‘Loden’ and S. schwerinii × S. viminalis cv. ‘Tora’) compared to plots with pure genotypes in a randomized block design in a field experiment in Northern Germany. We assessed the abundance of mycorrhizal colonization, fungal diversity, fine root density in the soil and activities of hydrolytic enzymes involved in P-mobilization in the mycorrhizosphere in autumn and following spring after three vegetation periods. Mycorrhizal and endophytic diversity was low under all Salix treatments with Laccaria tortilis being the dominating ectomyorrhizal fungal species, and Cadophora and Paraphaeosphaeria spp. being the most common endophytic fungi. Interspecific root competition increased richness and root colonization by endophytic fungi (four taxa in the mixture vs. one found in the pure host genotype cultures) more than by ectomycorrhizal fungi and increased the activities of hydrolytic soil enzymes involved in the P-mineralization (acid phosphatase and β-glucosidase) in mixed stands. The data suggest selective promotion of endophytic root colonization and changed competition for nutrients by mixture of Salix genotypes

Image_1_Mixture of Salix Genotypes Promotes Root Colonization With Dark Septate Endophytes and Changes P Cycling in the Mycorrhizosphere.pdf

<p>The roots of Salix spp. can be colonized by two types of mycorrhizal fungi (ectomycorrhizal and arbuscular) and furthermore by dark-septate endophytes. The fungal root colonization is affected by the plant genotype, soil properties and their interactions. However, the impact of host diversity accomplished by mixing different Salix genotypes within the site on root-associated fungi and P-mobilization in the field is not known. It can be hypothesized that mixing of genotypes with strong eco-physiological differences changes the diversity and abundance of root-associated fungi and P-mobilization in the mycorrhizosphere based on different root characteristics. To test this hypothesis, we have studied the mixture of two fundamentally eco-physiologically different Salix genotypes (S. dasyclados cv. ‘Loden’ and S. schwerinii × S. viminalis cv. ‘Tora’) compared to plots with pure genotypes in a randomized block design in a field experiment in Northern Germany. We assessed the abundance of mycorrhizal colonization, fungal diversity, fine root density in the soil and activities of hydrolytic enzymes involved in P-mobilization in the mycorrhizosphere in autumn and following spring after three vegetation periods. Mycorrhizal and endophytic diversity was low under all Salix treatments with Laccaria tortilis being the dominating ectomyorrhizal fungal species, and Cadophora and Paraphaeosphaeria spp. being the most common endophytic fungi. Interspecific root competition increased richness and root colonization by endophytic fungi (four taxa in the mixture vs. one found in the pure host genotype cultures) more than by ectomycorrhizal fungi and increased the activities of hydrolytic soil enzymes involved in the P-mineralization (acid phosphatase and β-glucosidase) in mixed stands. The data suggest selective promotion of endophytic root colonization and changed competition for nutrients by mixture of Salix genotypes.</p

Erythropoietin Restores Long-Term Neurocognitive Function Involving Mechanisms of Neuronal Plasticity in a Model of Hyperoxia-Induced Preterm Brain Injury

Cerebral white and grey matter injury is the leading cause of an adverse neurodevelopmental outcome in prematurely born infants. High oxygen concentrations have been shown to contribute to the pathogenesis of neonatal brain damage. Here, we focused on motor-cognitive outcome up to the adolescent and adult age in an experimental model of preterm brain injury. In search of the putative mechanisms of action we evaluated oligodendrocyte degeneration, myelination, and modulation of synaptic plasticity-related molecules. A single dose of erythropoietin (20,000 IU/kg) at the onset of hyperoxia (24 hours, 80% oxygen) in 6-day-old Wistar rats improved long-lasting neurocognitive development up to the adolescent and adult stage. Analysis of white matter structures revealed a reduction of acute oligodendrocyte degeneration. However, erythropoietin did not influence hypomyelination occurring a few days after injury or long-term microstructural white matter abnormalities detected in adult animals. Erythropoietin administration reverted hyperoxia-induced reduction of neuronal plasticity-related mRNA expression up to four months after injury. Thus, our findings highlight the importance of erythropoietin as a neuroregenerative treatment option in neonatal brain injury, leading to improved memory function in adolescent and adult rats which may be linked to increased neuronal network connectivity